Geographical subdivision,demographic history and gene flow in two sympatric species of intertidal snails, <Emphasis Type="Italic">Nerita scabricosta</Emphasis> and <Emphasis Type="Italic">Nerita funiculata</Emphasis>, from the tropical eastern Pacific

The patchy distribution of rocky intertidal communities in the tropical eastern Pacific (TEP) may impose severe constraints on the genetic connectivity among populations of marine invertebrates associated with this habitat. In this study, we analyzed a portion of the mitochondrial cytochrome c oxidase subunit I (COI) gene in two sympatric species of marine snails, Nerita scabricosta and Nerita funiculata, common inhabitants of the rocky intertidal from the Gulf of California (Sea of Cortez) and outer Pacific coast of the southern Baja California (Baja) peninsula to northern South America, to assess genetic connectivity among populations of each species. One of our aims was to determine whether the morphological, behavioral, and ecological differences observed among populations of both species throughout their range in the TEP corresponded to population genetic differences. In addition, we were interested in elucidating the demographic history of both species. We found no evidence of genetic structure throughout the Gulf of California and outer coast of the Baja peninsula region for either species. Comparisons between Gulf of California/Baja and Panama populations, however, showed significant genetic differentiation for N. scabricosta, but not for N. funiculata. The genetic differences between Mexican and Panamanian populations of N. scabricosta were consistent with previously reported ecological and behavioral differences for this species between these two distant regions. However, previously reported size differences between northern and central/southern Gulf of California individuals of N. scabricosta do not correspond with our findings of genetic connectivity among these populations. Results from neutrality tests (Tajima’s D and Fu’s F _S), the mismatch distribution, and Bayesian skyline analyses suggested that both species have experienced dramatic population expansions dating to the Pleistocene.     

Population genetic structure of escolar (<Emphasis Type="Italic">Lepidocybium flavobrunneum</Emphasis>)

Escolar (Lepidocybium flavobrunneum) is a large, mesopelagic fish that inhabits tropical and temperate seas throughout the world, and is a common bycatch in pelagic longline fisheries that target tuna and swordfish. Few studies have explored the biology and natural history of escolar, and little is known regarding its population structure. To evaluate the genetic basis of population structure of escolar throughout their range, we surveyed genetic variation over an 806 base pair fragment of the mitochondrial control region. In total, 225 individuals from six geographically distant locations throughout the Atlantic (Gulf of Mexico, Brazil, South Africa) and Pacific (Ecuador, Hawaii, Australia) were analyzed. A neighbor-joining tree of haplotypes based on maximum likelihood distances revealed two highly divergent clades (δ = 4.85%) that were predominantly restricted to the Atlantic and Indo-Pacific ocean basins. All Atlantic clade individuals occurred in the Atlantic Ocean and all but four Pacific clade individuals were found in the Pacific Ocean. The four Atlantic escolar with Pacific clade haplotypes were found in the South Africa collection. The nuclear ITS-1 gene region of these four individuals was subsequently analyzed and compared to the ITS-1 gene region of four individuals from the South Africa collection with Atlantic clade haplotypes as well as four representative individuals each from the Atlantic and Pacific collections. The four South Africa escolar with Pacific mitochondrial control region haplotypes all had ITS-1 gene region sequences that clustered with the Pacific escolar, suggesting that they were recent migrants from the Indo-Pacific. Due to the high divergence and geographic separation of the Atlantic and Pacific clades, as well as reported morphological differences between Atlantic and Indo-Pacific specimens, consideration of the Atlantic and Indo-Pacific populations as separate species or subspecies may be warranted, though further study is necessary.     

Genetic analysis of sympatric morphotypes of common dolphins (genus Delphinus)

Sympatric populations of two forms of the common dolphin, currently recognized collectively as Delphinus delphis Linnaeus, occur in several areas of the world's oceans. A molecular genetic study was initiated to determine whether these forms are genetically distinct in the Northeast Pacific. We compared mitochondrial DNA sequences from the control region and cytochrome b gene between specimens of the long-beaked and the short-beaked morphotypes collected between 1986 and 1989 off the coast of southern California. Additional short-beaked specimens collected from the eastern tropical Pacific (in 1978 and 1982) and the Black Sea (in 1989) were also compared. There were no shared mitochondrial DNA haplotypes between the two morphotypes, and both gene regions exhibited frequency and fixed nucleotide substitutions between the two morphotypes. This genetic differentiation, coupled with unique morphological characters of the short-beaked and long-beaked morphotypes determined in a parallel study, indicate that although sympatric, these populations of common dolphin are reproductively isolated from one another and may represent separate species.     

Population genetic structure of North American <Emphasis Type="Italic">Ophiactis</Emphasis> spp. brittle stars possessing hemoglobin

With the discovery of previously unreported populations of hemoglobin-possessing Ophiactis from the Texas coast in the Gulf of Mexico, an investigation into its population structure, including populations of O. simplex from the Pacific coast of California and O. rubropoda from the Atlantic coast of Florida, was undertaken using DNA sequence data from the mitochondrial COI gene. The reconstructed haplotype network suggests that California populations contain the ancestral source of mtDNA variation, and there is no evidence of recent introductions into Texas. Population genetic analyses reveal the California, Florida, and Texas Ophiactis populations to each be significantly differentiated from one another. Sequence divergence among the three areas is shallower than would be predicted given biogeographic history. Texas and Florida populations are equally genetically diverged from California populations as they are to one another, despite the greater potential for gene flow between these areas. The genetic distinctiveness of the Texas populations and the concordance of this pattern with phylogeographic patterns in other brittle star systems indicate an isolated and independent evolutionary history and we hypothesize that the three geographic regions included in this study each serve as hypotheses of population-level lineages that remain to be tested with independent sources of data.     

The relationship between population genetic structure and pelagic larval duration in coral reef fishes on the Great Barrier Reef

Pelagic larval duration (PLD) is a commonly used proxy for dispersal potential in coral reef fishes. Here we examine the relationship between PLD, genetic structure and genetic variability in geographically widespread and ecological generalist species from one coral reef fish family (Pomacentridae) that differs in mean larval duration by more than a month. The genetic structure was estimated in eight species using a mitochondrial molecular marker (D-loop) and in a sub-set of five species using nuclear molecular markers (ISSRs). Estimates of genetic differentiation were similar among species with pelagic larvae, but differed between molecular markers. The mtDNA indicated no structure in all species except one, while the ISSR indicated some structure between the sampling locations in all species. We detected a relationship between PLD and genetic structure using both markers. These relationships, however, were caused by a single species, Acanthochromis polyacanthus, which differs from all the other species examined here in lacking a larval phase. With this species excluded, there was no relationship between PLD and genetic structure using either marker despite a range of PLDs of more than 20 days. Genetic diversities were generally high in all species and did not differ significantly among species and locations. Nucleotide diversity and total heterozygosity were negatively related to maximum PLD but again these relationships were caused by A. polyacanthus and disappeared when this species was excluded. These genetic patterns are consistent with moderate gene flow among well-connected locations and indicate that at this phylogenetic level (i.e., within family) the duration of the pelagic larval phase is unrelated to the patterns of genetic differentiation.     

Sympatric size variants of the microcopepod Oncaea venusta exhibit distinct lineages in DNA sequences

We investigated the phylogenetic relationships among different size groups of the pelagic microcopepod Oncaea venusta Philippi, 1843, by comparing the patterns of genetic variation of specimens collected at five locations of the Indo-West Pacific Ocean. Phylogenetic analyses were based on sequence data obtained from two DNA markers: A 310 bp fragment of the mitochondrial cytochrome b (cyt b) gene and a 480 bp fragment of the nuclear internal transcribed spacer 1 (ITS1). The cyt b sequences showed a much higher level of variation than those from ITS1, but the conclusion from both genes was concordant. Four genetic clades could be differentiated. A small- and a large-size group were unambiguously assigned to two distinct clades or lineages. Unexpectedly, the medium-sized individuals could be divided into another two different genetic clades. All four lineages were supported by high bootstrap values. The high levels of sequence divergence under sympatric conditions indicated that at least the two main groups, the large and the small one, may be assigned to different species. For the medium-size group additional morphological studies and more sensitive nuclear markers are required to clarify their taxonomic status.     

Genetic structuring of <Emphasis Type="Italic">Latris lineata</Emphasis> at localized and transoceanic scales

Striped trumpeter (Latris lineata) is a demersal teleost distributed around the temperate clines of all the major oceans in the southern hemisphere. Within Tasmanian waters the species is managed as a single stock, although no studies have been performed to confirm genetic panmixia. A protracted pelagic larval phase and a recent transoceanic tag recapture of an adult fish suggest significant potential for genetic mixing between widely separated populations. Phylogenetic analysis of mitochondrial DNA control region sequences suggested no genetic mixing between Tasmania, New Zealand and St Paul/Amsterdam Islands, evidence for the first time that there is population structure at a transoceanic scale for this species. In addition, an analysis of molecular variance coupled with phylogenetic analyses suggested no significant structuring of striped trumpeter from three locations around Tasmania. The information provided in this study is useful for the design of modern fisheries management techniques such as spatially implemented marine reserves. In addition, species-by-species knowledge about population structures of marine species facilitates ecologically useful generalizations concerning their population dynamics and key issues on the broader ecology of the oceans.     

Phylogeography of two Atlantic squirrelfishes (Family Holocentridae): exploring links between pelagic larval duration and population connectivity

Genetic surveys of reef fishes have revealed high population connectivity within ocean basins, consistent with the assumption that pelagic larvae disperse long distances by oceanic currents. However, several recent studies have demonstrated that larval retention and self-recruitment may be higher than previously expected. To assess connectivity in tropical reef fishes, we contribute range-wide mtDNA surveys of two Atlantic squirrelfishes (family Holocentridae). The blackbar soldierfish, Myripristis jacobus, has a pelagic juvenile phase of about 58 days, compared to about 71 days (~22% longer) in the longjaw squirrelfish, Holocentrus ascensionis. If the pelagic duration is guiding dispersal ability, M. jacobus should have greater population genetic structure than H. ascensionis. In comparisons of mtDNA cytochrome b sequences from 69 M. jacobus (744 bp) and 101 H. ascensionis (769 bp), both species exhibited a large number of closely related haplotypes (h=0.781 and 0.974, π=0.003 and 0.006, respectively), indicating late Pleistocene coalescence of mtDNA lineages. Contrary to the prediction based on pelagic duration, M. jacobus has much less population structure (φ_ST=0.008, P=0.228) than H. ascensionis (φ_ST=0.091, P<0.001). Significant population partitions in H. ascensionis were observed between eastern, central and western Atlantic, and between Brazil and the Caribbean in the western Atlantic. These results, in combination with the findings from 13 codistributed species, indicate that pelagic larval duration is a poor predictor of population genetic structure in Atlantic reef fishes. A key to understanding this disparity may be the evolutionary depth among corresponding taxonomic groups of “reef fishes”, which extends back to the mid-Cretaceous and encompasses enormous diversity in ecology and life history. We should not expect a simple relationship between pelagic larval duration and genetic connectivity, among lineages that diverged 50–100 million years ago.     

MtDNA population structure and gene flow in lingcod (<Emphasis Type="Italic">Ophiodon elongatus</Emphasis>): limited connectivity despite long-lived pelagic larvae

Lingcod, Ophiodon elongatus Girard, have a 3-month pelagic larval stage and are an important recreational and commercial species on the west coast of North America. Cytochrome-c oxidase I sequences from tissue samples were used to characterize population structure and infer patterns of gene flow from California to Alaska. No significant genetic structure was found when estimates of Wright’s F _ST (i.e., Φ_ST) were generated among all populations sampled. Nesting populations within regions, however, indicated that the inner coast of Washington State is distinct, a result corroborating previous allozyme work. Coalescent-based estimates of gene flow indicate that although migration can be high from an evolutionary perspective, nearly half of all comparisons among populations showed no gene flow in at least one direction. From an ecological perspective, moderate migration rates (Nm < 10) among most populations provide surprisingly limited connectivity at large (∼ 1,000 km) and small (∼100 km) spatial scales. Coalescent-based estimates also show that gene flow between the inner and the outer coasts is asymmetric, a result consistent with prevailing surface currents. Because the expected inter-locus variances for coalescent-based estimates of gene flow are likely large, future work will benefit from analyses of nuclear DNA markers. However, limited demographic connectivity on large spatial scales may help explain why stock recovery has been uneven, with greater recovery in the northern (87% rebuilt) than in the southern (24% rebuilt) fishery region, supporting a regional management strategy. These results suggest that despite a 3-month pelagic larval stage, some areas may be effectively closed with respect to both population dynamics and fishery management issues.     

Phylogeography, historical demography, and the role of post-settlement ecology in two Hawaiian damselfish species

Coral reef fish generally have relatively sedentary juvenile and adult phases and a presumed highly dispersive pelagic larval phase, yet previous studies that have tried to relate pelagic larval duration (PLD) to population structure have given inconsistent results. In the present study, the population structures of two damselfishes, Stegastes fasciolatus and Dascyllus albisella, were examined using mitochondrial control region sequences. The two species have similar PLDs (∼25 and 27 days respectively), but consistently differ in their settlement preferences, habitat, and densities in populations throughout the Hawaiian Archipelago, from Hawaii north to Kure Atoll, and south to Johnston Atoll. Information on habitat preferences and population densities were collected between September 2000 and October 2002, and tissue samples for the genetic studies were collected between January and April 2004. Based on the differences in habitat and abundance of the two species, the expectation was that S. fasciolatus would have high genetic variability but little population structure compared to D. albisella, and this was largely confirmed. Stegastes fasciolatus had little population structure in most of the Hawaiian Islands, and D. albisella showed evidence of strong population structure throughout its range. An exception to this pattern was the large difference between the Kure Atoll population of S. fasciolatus and all others. These results suggest that the interaction of several biological factors (e.g. species-specific spawning habitat and season) with environmental factors (e.g. seasonal wind and current patterns) may have more influence on population structure than single life history characteristics, such as the PLD.     

Cryptic isolation of Gulf of California shovelnose guitarfish evidenced by mitochondrial DNA

The shovelnose guitarfish Rhinobatos productus is an evolutionarily, ecologically, and economically important ray, with a continuous distribution from San Francisco, California (USA), to Mazatlan, Sinaloa, and in the Gulf of California (Mexico). Regional studies have revealed morphometric differences between shovelnose from the Gulf of California and the Pacific coast of Baja California, which may result from phenotypic plasticity in the presence of high levels of gene flow or from a degree of genetic differentiation in the presence of cryptic isolation within a continuous distribution. We used PCR-RFLP of the mitochondrial control region to assess the degree of genetic differentiation between Gulf of California and Pacific shovelnose guitarfish. We found very high levels of molecular diversity (averages: h=0.77, =1.19%), which may be associated with historically large and stable populations, as well as very significant levels of genetic differentiation between gulf and Pacific samples (²=64, P<0.0001; _ST=0.63, P<0.0001, mean nucleotide divergence d=2.47%). We found a deep phylogeographic break between haplotypes from the gulf and the Pacific, which may suggest the existence of cryptic species but clearly indicates more than one evolutionarily significant unit of R. productus. Our results show a pattern of genetic structure and levels of differentiation consistent with the geological history of the region. Furthermore, these findings have wide-ranging implications for the management and conservation of cartilaginous fish in Mexico, as they reveal the existence of biological diversity that will go unnoticed without the genetic scrutiny of intraspecific variation and that is highly relevant for much needed management and conservation efforts.Communicated by P.W. Sammarco, Chauvin     

Nuclear and mtDNA lineage diversity in wild and cultured Pacific lion-paw scallop, Nodipecten subnodosus (Baja California Peninsula, Mexico)

Pacific lion-paw scallops were collected from natural aggregations in Laguna Ojo de Liebre (Pacific Ocean), the Gulf of California, and from aquaculture facilities for genetic diversity analyses. Mitochondrial DNA sequencing uncovered two highly supported clades separated by 2.5% divergence. Data from ten microsatellite markers suggest individuals from these mitogroups are introgressed, raising questions about the mitotype origin. Some evidence suggests gene flow between La Paz and Ojo de Liebre; otherwise the Gulf of California and Ojo de Liebre are acting as two distinct populations. It is unclear whether translocations between sites have influenced the observed genetic structure or whether gene flow has been facilitated by past geologic events. Finally, scallops spawned for aquaculture are unique from the wild and have significantly less diversity. These results warrant the attention of managers and producers who should work to monitor and conserve genetic diversity in both wild and aquaculture populations.     

Spatial overlap and trophic interactions between pelagic fish and large jellyfish in the northern California Current

Recent studies have indicated that populations of gelatinous zooplankton may be increasing and expanding in geographic coverage, and these increases may in turn affect coastal fish populations. We conducted trawl surveys in the northern California Current and documented a substantial biomass of scyphomedusae consisting primarily of two species (Chrysaora fuscescens and Aurelia labiata). Spatial overlap of these jellyfish with most pelagic fishes, including salmon, was generally low, but there were regions of relatively high overlap where trophic interactions may have been occurring. We compared feeding ecology of jellyfish and pelagic fishes based on diet composition and found that trophic overlap was high with planktivorous species that consume copepods and euphausiid eggs such as Pacific sardines (Sardinops sagax), northern anchovy (Engraulis mordax), Pacific saury (Cololabis saira), and Pacific herring (Clupea pallasi). Moreover, isotope and diet analyses suggest that jellyfish occupy a trophic level similar to that of small pelagic fishes such as herring, sardines and northern anchovy. Thus jellyfish have the potential, given their substantial biomass, of competing with these species, especially in years with low ecosystem productivity where prey resources will be limited.     

Diversity of free-living marine nematodes (Enoplida) from Baja California assessed by integrative taxonomy

We used morphological and molecular approaches to evaluate the diversity of free-living marine nematodes (order Enoplida) at four coastal sites in the Gulf of California and three on the Pacific coast of Baja California, Mexico. We identified 22 morphological species belonging to six families, of which Thoracostomopsidae and Oncholaimidae were the most diverse. The genus Mesacanthion (Thoracostomopsidae) was the most widespread and diverse. Five allopatric species, genetically and morphologically differentiated, were found in two localities in the Gulf of California (M. sp1 and M. sp2) and three in the Pacific coast (M. sp3, M. sp4 and M. sp5). Overall, we produced 19 and 20 sequences for the 18S and 28S genes, respectively. Neither gene displayed intraspecific polymorphisms, which allowed us to establish that some morphological variation was likely either ontogenetic or due to phenotypic plasticity. Although 18S and 28S phylogenies were topologically congruent (incongruence length difference test, P > 0.05), divergences between species were much higher in the 28S gene. Moreover, this gene possessed a stronger phylogenetic signal to resolve relationships involving Rhabdodemania and Bathylaimus. On the other hand, the close relationship of Pareurystomina (Enchilidiidae) with oncholaimids warrants further study. The 28S sequences (D2D3 domain) may be better suited for DNA barcoding of marine nematodes than those from the 18S rDNA, particularly for differentiating closely related or cryptic species. Finally, our results underline the relevance of adopting an integrative approach encompassing morphological and molecular analyses to improve the assessment of marine nematode diversity and advance their taxonomy.     

Population divergence in the sinistral whelks of North America,with special reference to the east Florida ecotone

This study evaluated models of species relationships among sinistral whelks in the genus Busycon in the Atlantic and Gulf of Mexico. Gene frequencies at eight polymorphic allozyme loci, shell morphology, anatomy, and partial DNA sequences for the cytochrome c oxidase I (COI) mitochondrial gene were examined in eight populations, ranging from New Jersey to the Yucatan peninsula, and from the dextrally coiled sister taxon Busycon carica (Gmelin, 1791). Whelks were collected in 1997 and 1998. The maximum COI sequence divergence recorded among 32 sinistral individuals was 1.96%, which together with the absence of any gross or qualitative morphological differences, suggested all eight populations should be considered conspecific. High levels of divergence between the allopatric western Atlantic and Gulf of Mexico populations, as revealed by fixed or nearly fixed differences at several allozyme-encoding loci were interpreted as evidence that the east Florida ecotone constitutes a significant barrier to gene flow. Size trimming also revealed several significant quantitative differences in shell and radular morphology between the three pooled Atlantic populations and five pooled Gulf populations. The Yucatan sample was the most distinctive conchologically, with heavy spines and tumid ridges, possibly related to stone crab predation. Based on the evidence all left-handed whelks of North America should be referred to the oldest available nomen, Busycon perversum (Linné, 1758), with three subspecies, B. perversum perversum along the Yucatan peninsula, B. perversum sinistrum (Hollister, 1958) in the northern and eastern Gulf of Mexico, and B. perversum laeostomum (Kent, 1982) in the Atlantic.Communicated by J.P. Grassle, New Brunswick     

Structure and migration corridors in Pacific populations of the Swordfish Xiphius gladius, as inferred through analyses of mitochondrial DNA

 The swordfish Xiphias gladius is a migratory oceanic species distributed in sub-tropical and temperate waters worldwide. Studies utilizing mitochondrial DNA (mtDNA) have demonstrated genetic subdivision between ocean basins, as well as within the Atlantic basin. However, there has been no support of population subdivision within the Pacific. We sequenced 629 base pairs of the control region for 281 swordfish collected in the Pacific. A rate heterogeneity parameter, alpha, was found to be 0.201, indicating substantial variation in mutation rate within the control region of swordfish. Hierarchical analysis of molecular variance supported significant genetic structuring among Pacific populations. Northern and southern populations in the western Pacific were significantly divergent, while populations in the east appeared to be genetically continuous. Regression analysis supported a correlation of genetic differentiation with geographic distance along a U-shaped corridor of gene flow. These results reveal a pelagic biogeographic pattern heretofore unrecognized in the Pacific, and reject the null hypothesis that Pacific populations of swordfish are unstructured and comprise only a single homogeneous stock. Received: 10 November 1998 / Accepted: 4 February 2000     

Population structure of the planktonic copepod Calanus pacificus in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Gene flow at three spatial scales in a coral reef fish, the three-spot dascyllus, Dascyllus trimaculatus

Dispersal in coral reef fishes occurs predominantly during the larval planktonic stage of their life cycle. With relatively brief larval stages, damselfishes (Pomacentridae) are likely to exhibit limited dispersal. This study evaluates gene flow at three spatial scales in one species of coral reef damselfish, Dascyllus trimaculatus. Samples were collected at seven locations at Moorea, Society Islands, French Polynesia. Phylogenetic relationships and gene flow based on mitochondrial control region DNA sequences between these locations were evaluated (first spatial scale). Although spatial structure was not found, molecular markers showed clear temporal structure, which may be because pulses of settling larvae have distinct genetic composition. Moorea samples were then compared with individuals from a distant island (750 km), Rangiroa, Tuamotu Archipelago, French Polynesia (second spatial scale). Post-recruitment events (selection) and gene flow were probably responsible for the lack of structure observed between populations from Moorea and Rangiroa. Finally, samples from six Indo-West Pacific locations, Zanzibar, Indonesia, Japan, Christmas Island, Hawaii, and French Polynesia were compared (third spatial scale). Strong population structure was observed between Indo-West Pacific populations. Received: 26 May 2000 / Accepted: 10 October 2000     

Are meiofaunal species cosmopolitan? Morphological and molecular analysis of Xenotrichula intermedia (Gastrotricha: Chaetonotida)

Many meiofaunal species are reported to be cosmopolitan, but due to uncertainties of identification, the affiliation of specimens from geographically distant areas to the same species-taxon is problematic. In this study, we examined morphological and molecular variation in samples of Xenotrichula intermedia Remane (Gastrotricha: Chaetonotida) from the Mediterranean Sea, the northwestern Atlantic and the northern Gulf of Mexico. Univariate analysis of 16 morphological traits was unable to detect differences among populations, except for the length of the pharynx, which was significantly shorter in the Gulf of Mexico specimens. Canonical discriminant analysis separated the Gulf of Mexico specimens from the other two populations, with pharynx length contributing about half of the total discrimination. Molecular analysis based on restriction-fragment length polymorphisms (RFLPs) in a 710-base pair polymerase chain-reaction (PCR) produet representing roughly half of the mitochondrial cytochrome oxidase I (COI) gene detected four haplotypes: one each from the Mediterranean and the Gulf of Mexico populations and two coexisting within the Atlantic population. The estimated nucleotide-sequence divergence calculated for each pairwise combination of haplotypes (based on the proportion of shared fragments) ranged from 5.3 to 11.5%. The high genetic divergence and the inability to clearly separate populations based on morphology suggest that individuals characterized by different haplotypes are genetically isolated sibling species.     

Population structure of the planktonic copepod <Emphasis Type="Italic">Calanus pacificus</Emphasis> in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.